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class=\u0022elements-frag-data highwire-markup\u0022 id=\u0022fig-data\u0022\u003E\u003Cdiv id=\u0022fig-data-figures\u0022 class=\u0022group frag-figures\u0022\u003E\u003Cdiv class=\u0022fig-data-title-jump clearfix\u0022\u003E\u003Ch3 id=\u0022fig-frag-data-title\u0022 class=\u0022fig-data-group-title\u0022\u003EFigures\u003C\/h3\u003E\u003Cdiv class=\u0022fig-data-jump-links\u0022\u003E\u003C\/div\u003E\u003C\/div\u003E\u003Cdiv class=\u0022item-list\u0022\u003E\u003Cul id=\u0022fig-frag-fig\u0022 class=\u0022fig-frag-data-list clearfix\u0022\u003E\u003Cli class=\u0022first\u0022\u003E\u003Cdiv class=\u0022element-fig-frag-data clearfix supplementary-material-caption\u0022\u003E\u003Cdiv class=\u0022highwire-markup\u0022\u003E\u003Cdiv xmlns=\u0022http:\/\/www.w3.org\/1999\/xhtml\u0022 id=\u0022content-block-markup\u0022 xmlns:xhtml=\u0022http:\/\/www.w3.org\/1999\/xhtml\u0022\u003E\u003Cdiv class=\u0022fig-expansion\u0022 id=\u0022F1\u0022\u003E\u003Cspan class=\u0022highwire-journal-article-marker-start\u0022\u003E\u003C\/span\u003E\u003Cdiv class=\u0022highwire-figure\u0022\u003E\u003Cdiv class=\u0022fig-inline-img-wrapper\u0022\u003E\u003Cdiv class=\u0022fig-inline-img\u0022\u003E\u003Ca href=\u0022http:\/\/jcs.biologists.org\/content\/joces\/118\/10\/2341\/F1.large.jpg?width=800\u0026amp;height=600\u0026amp;carousel=1\u0022 title=\u0022Poleward MT flux in a metaphase-stage mitotic spindle. Flux occurs on both kMTs (red lines) and non-kMTs (blue lines) in the spindle. Tubulin subunits are incorporated into polymer at MT plus-ends and removed at their minus-ends focused at the spindle poles. Arrows within the red and blue lines indicate the direction of continuous ATP-dependent polymer movement. kMT plus-end assembly stops at the transition to anaphase (although there are exceptions to this rule; Chen and Zhang, 2004; LaFountain et al., 2004; see text). Astral MTs (green lines), whose minus-ends are embedded in the centrosomes, do not flux. Orange arrows above the spindle indicate the poleward direction of force exerted by flux on each sister kinetochore. Likewise, opposing blue arrows indicate the metaphase plateward direction of force exerted by flux on each spindle pole. Importantly, the major source of MT assembly dynamics in the spindle is plus-end dynamic instability, which is not shown here for simplicity.\u0022 class=\u0022highwire-fragment fragment-images colorbox-load\u0022 rel=\u0022gallery-fragment-images-1492496706\u0022 data-figure-caption=\u0022\u0026lt;div class=\u0026quot;highwire-markup\u0026quot;\u0026gt;Poleward MT flux in a metaphase-stage mitotic spindle. Flux occurs on both kMTs (red lines) and non-kMTs (blue lines) in the spindle. Tubulin subunits are incorporated into polymer at MT plus-ends and removed at their minus-ends focused at the spindle poles. Arrows within the red and blue lines indicate the direction of continuous ATP-dependent polymer movement. kMT plus-end assembly stops at the transition to anaphase (although there are exceptions to this rule; Chen and Zhang, 2004; LaFountain et al., 2004; see text). Astral MTs (green lines), whose minus-ends are embedded in the centrosomes, do not flux. Orange arrows above the spindle indicate the poleward direction of force exerted by flux on each sister kinetochore. Likewise, opposing blue arrows indicate the metaphase plateward direction of force exerted by flux on each spindle pole. Importantly, the major source of MT assembly dynamics in the spindle is plus-end dynamic instability, which is not shown here for simplicity.\u0026lt;\/div\u0026gt;\u0022 data-icon-position=\u0022\u0022 data-hide-link-title=\u00220\u0022\u003E\u003Cspan class=\u0022hw-responsive-img\u0022\u003E\u003Cimg class=\u0022highwire-fragment fragment-image lazyload\u0022 alt=\u0022 Fig. 1. \u0022 src=\u0022data:image\/gif;base64,R0lGODlhAQABAIAAAAAAAP\/\/\/yH5BAEAAAAALAAAAAABAAEAAAIBRAA7\u0022 data-src=\u0022http:\/\/jcs.biologists.org\/content\/joces\/118\/10\/2341\/F1.medium.gif\u0022\/\u003E\u003Cnoscript\u003E\u003Cimg class=\u0022highwire-fragment fragment-image\u0022 alt=\u0022 Fig. 1. \u0022 src=\u0022http:\/\/jcs.biologists.org\/content\/joces\/118\/10\/2341\/F1.medium.gif\u0022\/\u003E\u003C\/noscript\u003E\u003C\/span\u003E\u003C\/a\u003E\u003C\/div\u003E\u003C\/div\u003E\u003Cul class=\u0022highwire-figure-links inline\u0022\u003E\u003Cli class=\u0022download-fig first\u0022\u003E\u003Ca href=\u0022http:\/\/jcs.biologists.org\/content\/joces\/118\/10\/2341\/F1.large.jpg?download=true\u0022 class=\u0022highwire-figure-link highwire-figure-link-download\u0022 title=\u0022Download Fig. 1. \u0022 data-icon-position=\u0022\u0022 data-hide-link-title=\u00220\u0022\u003EDownload figure\u003C\/a\u003E\u003C\/li\u003E\u003Cli class=\u0022new-tab\u0022\u003E\u003Ca href=\u0022http:\/\/jcs.biologists.org\/content\/joces\/118\/10\/2341\/F1.large.jpg\u0022 class=\u0022highwire-figure-link highwire-figure-link-newtab\u0022 target=\u0022_blank\u0022 data-icon-position=\u0022\u0022 data-hide-link-title=\u00220\u0022\u003EOpen in new tab\u003C\/a\u003E\u003C\/li\u003E\u003Cli class=\u0022download-ppt last\u0022\u003E\u003Ca href=\u0022\/highwire\/powerpoint\/1564453\u0022 class=\u0022highwire-figure-link highwire-figure-link-ppt\u0022 data-icon-position=\u0022\u0022 data-hide-link-title=\u00220\u0022\u003EDownload powerpoint\u003C\/a\u003E\u003C\/li\u003E\u003C\/ul\u003E\u003C\/div\u003E\u003Cdiv class=\u0022fig-caption\u0022 xmlns:xhtml=\u0022http:\/\/www.w3.org\/1999\/xhtml\u0022\u003E\u003Cspan class=\u0022fig-label\u0022\u003E\n \u003Cstrong\u003EFig. 1.\u003C\/strong\u003E\n \u003C\/span\u003E \n \u003Cp id=\u0022p-5\u0022\u003EPoleward MT flux in a metaphase-stage mitotic spindle. Flux occurs on both kMTs (red lines) and non-kMTs (blue lines) in the spindle. Tubulin subunits are incorporated into polymer at MT plus-ends and removed at their minus-ends focused at the spindle poles. Arrows within the red and blue lines indicate the direction of continuous ATP-dependent polymer movement. kMT plus-end assembly stops at the transition to anaphase (although there are exceptions to this rule; Chen and Zhang, 2004; LaFountain et al., 2004; see text). Astral MTs (green lines), whose minus-ends are embedded in the centrosomes, do not flux. Orange arrows above the spindle indicate the poleward direction of force exerted by flux on each sister kinetochore. Likewise, opposing blue arrows indicate the metaphase plateward direction of force exerted by flux on each spindle pole. Importantly, the major source of MT assembly dynamics in the spindle is plus-end dynamic instability, which is not shown here for simplicity.\u003C\/p\u003E\n \u003Cdiv class=\u0022sb-div caption-clear\u0022\u003E\u003C\/div\u003E\u003C\/div\u003E\u003Cspan class=\u0022highwire-journal-article-marker-end\u0022\u003E\u003C\/span\u003E\u003C\/div\u003E\u003Cspan id=\u0022related-urls\u0022\u003E\u003C\/span\u003E\u003C\/div\u003E\u003C\/div\u003E\u003C\/div\u003E\u003C\/li\u003E\u003Cli\u003E\u003Cdiv class=\u0022element-fig-frag-data clearfix supplementary-material-caption\u0022\u003E\u003Cdiv class=\u0022highwire-markup\u0022\u003E\u003Cdiv xmlns=\u0022http:\/\/www.w3.org\/1999\/xhtml\u0022 id=\u0022content-block-markup\u0022 xmlns:xhtml=\u0022http:\/\/www.w3.org\/1999\/xhtml\u0022\u003E\u003Cdiv class=\u0022fig-expansion\u0022 id=\u0022F2\u0022\u003E\u003Cspan class=\u0022highwire-journal-article-marker-start\u0022\u003E\u003C\/span\u003E\u003Cdiv class=\u0022highwire-figure\u0022\u003E\u003Cdiv class=\u0022fig-inline-img-wrapper\u0022\u003E\u003Cdiv class=\u0022fig-inline-img\u0022\u003E\u003Ca href=\u0022http:\/\/jcs.biologists.org\/content\/joces\/118\/10\/2341\/F2.large.jpg?width=800\u0026amp;height=600\u0026amp;carousel=1\u0022 title=\u0022A model for flux. (A) MT minus-end release from centrosomes could occur either by separation from the \u0026#x3B3;-tubulin ring complex (\u0026#x3B3;-TuRC) at or near the centrosome or from the MT-severing activity of centrosome-associated katanin (McNally et al., 1996). (B) A Kin I kinesin is targeted and tethered to an insoluble spindle pole matrix that anchors the spindle pole to the centrosome. Kin I actively drives flux (depicted as blue lines with arrows) by disassembling MT minus-ends. (C) This activity produces a polymer-free gap between the centrosome and the spindle pole that is observed in both live (top panel) and fixed (bottom panel) Drosophila syncytial blastoderm-stage embryos. The top panel shows rhodamine-labeled MTs (red) and GFP-histones (green). Indirect immunofluorescence in the bottom panel shows MTs (red) and KLP10A (green), which localizes within the gap and on centrosomes. (D) Poleward MT flux is driven in the metaphase half-spindle by a spindle-pole-associated Kin I kinesin. kMT length is maintained by the activity of the kinetochore-associated CLASP protein that induces plus-end polymerization.\u0022 class=\u0022highwire-fragment fragment-images colorbox-load\u0022 rel=\u0022gallery-fragment-images-1492496706\u0022 data-figure-caption=\u0022\u0026lt;div class=\u0026quot;highwire-markup\u0026quot;\u0026gt;A model for flux. (A) MT minus-end release from centrosomes could occur either by separation from the \u0026#x3B3;-tubulin ring complex (\u0026#x3B3;-TuRC) at or near the centrosome or from the MT-severing activity of centrosome-associated katanin (McNally et al., 1996). (B) A Kin I kinesin is targeted and tethered to an insoluble spindle pole matrix that anchors the spindle pole to the centrosome. Kin I actively drives flux (depicted as blue lines with arrows) by disassembling MT minus-ends. (C) This activity produces a polymer-free gap between the centrosome and the spindle pole that is observed in both live (top panel) and fixed (bottom panel) Drosophila syncytial blastoderm-stage embryos. The top panel shows rhodamine-labeled MTs (red) and GFP-histones (green). Indirect immunofluorescence in the bottom panel shows MTs (red) and KLP10A (green), which localizes within the gap and on centrosomes. (D) Poleward MT flux is driven in the metaphase half-spindle by a spindle-pole-associated Kin I kinesin. kMT length is maintained by the activity of the kinetochore-associated CLASP protein that induces plus-end polymerization.\u0026lt;\/div\u0026gt;\u0022 data-icon-position=\u0022\u0022 data-hide-link-title=\u00220\u0022\u003E\u003Cspan class=\u0022hw-responsive-img\u0022\u003E\u003Cimg class=\u0022highwire-fragment fragment-image lazyload\u0022 alt=\u0022 Fig. 2. \u0022 src=\u0022data:image\/gif;base64,R0lGODlhAQABAIAAAAAAAP\/\/\/yH5BAEAAAAALAAAAAABAAEAAAIBRAA7\u0022 data-src=\u0022http:\/\/jcs.biologists.org\/content\/joces\/118\/10\/2341\/F2.medium.gif\u0022\/\u003E\u003Cnoscript\u003E\u003Cimg class=\u0022highwire-fragment fragment-image\u0022 alt=\u0022 Fig. 2. \u0022 src=\u0022http:\/\/jcs.biologists.org\/content\/joces\/118\/10\/2341\/F2.medium.gif\u0022\/\u003E\u003C\/noscript\u003E\u003C\/span\u003E\u003C\/a\u003E\u003C\/div\u003E\u003C\/div\u003E\u003Cul class=\u0022highwire-figure-links inline\u0022\u003E\u003Cli class=\u0022download-fig first\u0022\u003E\u003Ca href=\u0022http:\/\/jcs.biologists.org\/content\/joces\/118\/10\/2341\/F2.large.jpg?download=true\u0022 class=\u0022highwire-figure-link highwire-figure-link-download\u0022 title=\u0022Download Fig. 2. \u0022 data-icon-position=\u0022\u0022 data-hide-link-title=\u00220\u0022\u003EDownload figure\u003C\/a\u003E\u003C\/li\u003E\u003Cli class=\u0022new-tab\u0022\u003E\u003Ca href=\u0022http:\/\/jcs.biologists.org\/content\/joces\/118\/10\/2341\/F2.large.jpg\u0022 class=\u0022highwire-figure-link highwire-figure-link-newtab\u0022 target=\u0022_blank\u0022 data-icon-position=\u0022\u0022 data-hide-link-title=\u00220\u0022\u003EOpen in new tab\u003C\/a\u003E\u003C\/li\u003E\u003Cli class=\u0022download-ppt last\u0022\u003E\u003Ca href=\u0022\/highwire\/powerpoint\/1564459\u0022 class=\u0022highwire-figure-link highwire-figure-link-ppt\u0022 data-icon-position=\u0022\u0022 data-hide-link-title=\u00220\u0022\u003EDownload powerpoint\u003C\/a\u003E\u003C\/li\u003E\u003C\/ul\u003E\u003C\/div\u003E\u003Cdiv class=\u0022fig-caption\u0022 xmlns:xhtml=\u0022http:\/\/www.w3.org\/1999\/xhtml\u0022\u003E\u003Cspan class=\u0022fig-label\u0022\u003E\n \u003Cstrong\u003EFig. 2.\u003C\/strong\u003E\n \u003C\/span\u003E \n \u003Cp id=\u0022p-7\u0022\u003EA model for flux. (A) MT minus-end release from centrosomes could occur either by separation from the \u03b3-tubulin ring complex (\u03b3-TuRC) at or near the centrosome or from the MT-severing activity of centrosome-associated katanin (McNally et al., 1996). (B) A Kin I kinesin is targeted and tethered to an insoluble spindle pole matrix that anchors the spindle pole to the centrosome. Kin I actively drives flux (depicted as blue lines with arrows) by disassembling MT minus-ends. (C) This activity produces a polymer-free gap between the centrosome and the spindle pole that is observed in both live (top panel) and fixed (bottom panel) \u003Cem\u003EDrosophila\u003C\/em\u003E syncytial blastoderm-stage embryos. The top panel shows rhodamine-labeled MTs (red) and GFP-histones (green). Indirect immunofluorescence in the bottom panel shows MTs (red) and KLP10A (green), which localizes within the gap and on centrosomes. (D) Poleward MT flux is driven in the metaphase half-spindle by a spindle-pole-associated Kin I kinesin. kMT length is maintained by the activity of the kinetochore-associated CLASP protein that induces plus-end polymerization.\u003C\/p\u003E\n \u003Cdiv class=\u0022sb-div caption-clear\u0022\u003E\u003C\/div\u003E\u003C\/div\u003E\u003Cspan class=\u0022highwire-journal-article-marker-end\u0022\u003E\u003C\/span\u003E\u003C\/div\u003E\u003Cspan id=\u0022related-urls\u0022\u003E\u003C\/span\u003E\u003C\/div\u003E\u003C\/div\u003E\u003C\/div\u003E\u003C\/li\u003E\u003Cli class=\u0022last\u0022\u003E\u003Cdiv class=\u0022element-fig-frag-data clearfix supplementary-material-caption\u0022\u003E\u003Cdiv class=\u0022highwire-markup\u0022\u003E\u003Cdiv xmlns=\u0022http:\/\/www.w3.org\/1999\/xhtml\u0022 id=\u0022content-block-markup\u0022 xmlns:xhtml=\u0022http:\/\/www.w3.org\/1999\/xhtml\u0022\u003E\u003Cdiv class=\u0022fig-expansion\u0022 id=\u0022F3\u0022\u003E\u003Cspan class=\u0022highwire-journal-article-marker-start\u0022\u003E\u003C\/span\u003E\u003Cdiv class=\u0022highwire-figure\u0022\u003E\u003Cdiv class=\u0022fig-inline-img-wrapper\u0022\u003E\u003Cdiv class=\u0022fig-inline-img\u0022\u003E\u003Ca href=\u0022http:\/\/jcs.biologists.org\/content\/joces\/118\/10\/2341\/F3.large.jpg?width=800\u0026amp;height=600\u0026amp;carousel=1\u0022 title=\u0022Mechanistic models for bivalent positioning and congression in meiotic spindles of grasshopper spermatocytes, using the force from flux and plus-end kMT assembly. (A) Bivalent positioning. A bivalent maintains an equilibrium position at the spindle equator owing to an equivalent amount of flux (blue arrows) on homologous kinetochores attached to an equal number of kMTs. Moderate tension on the kinetochores induces kMT plus-end assembly (red arrows). Laser irradiation (green line) partially destroys a kinetochore, which reduces the number of kMTs to which it can bind. This sudden imbalance in kMT number increases the stress on the remaining kMTs of the irradiated kinetochore, inducing a greater rate of kMT plus-end assembly (red arrow). Compressive force on the opposing kMTs inhibits or decreases the rate of plus-end assembly, and, consequently, the bivalent moves (orange arrow) towards the pole that has the greater number of kMT attachments and the larger flux-generated force (larger blue arrow). As chromosome arms bind to an increasing density of spindle MTs that resist their poleward movement (yellow arrows), the increased tension on the unirradiated kinetochore induces plus-end polymerization. A new equilibrium position is reached when tension-induced plus-end assembly on the unirradiated kinetochore equals the rate of disassembly at the pole. (B) Congression. Initially, a bivalent close to one spindle pole becomes mono-oriented and is pulled poleward by flux (blue arrow). Chromosome arms bind to an increasing density of spindle MTs and the resulting resistance (yellow arrows) increases kinetochore tension to induce plus-end polymerization (red arrows). Poleward movement stops, facilitating the capture of the homologous kinetochore by the opposite pole (blue arrows in the spindle). Capture of the unattached kinetochore produces an even greater amount of tension and polymerization (red arrow) at the opposite kinetochore, allowing the bi-oriented bivalent to move to the spindle equator (orange arrow), even though kMT number and flux-generated force (blue arrows) are greater at the lagging homologous kinetochore. Finally, an equilibrium position is established at the spindle equator when the leading kinetochore is captured by an equal number of kMTs. Poleward force from flux (blue arrows) is equivalent in each half-spindle and is balanced by an equal rate of kMT plus-end assembly (red arrows) induced by moderate tension.\u0022 class=\u0022highwire-fragment fragment-images colorbox-load\u0022 rel=\u0022gallery-fragment-images-1492496706\u0022 data-figure-caption=\u0022\u0026lt;div class=\u0026quot;highwire-markup\u0026quot;\u0026gt;Mechanistic models for bivalent positioning and congression in meiotic spindles of grasshopper spermatocytes, using the force from flux and plus-end kMT assembly. (A) Bivalent positioning. A bivalent maintains an equilibrium position at the spindle equator owing to an equivalent amount of flux (blue arrows) on homologous kinetochores attached to an equal number of kMTs. Moderate tension on the kinetochores induces kMT plus-end assembly (red arrows). Laser irradiation (green line) partially destroys a kinetochore, which reduces the number of kMTs to which it can bind. This sudden imbalance in kMT number increases the stress on the remaining kMTs of the irradiated kinetochore, inducing a greater rate of kMT plus-end assembly (red arrow). Compressive force on the opposing kMTs inhibits or decreases the rate of plus-end assembly, and, consequently, the bivalent moves (orange arrow) towards the pole that has the greater number of kMT attachments and the larger flux-generated force (larger blue arrow). As chromosome arms bind to an increasing density of spindle MTs that resist their poleward movement (yellow arrows), the increased tension on the unirradiated kinetochore induces plus-end polymerization. A new equilibrium position is reached when tension-induced plus-end assembly on the unirradiated kinetochore equals the rate of disassembly at the pole. (B) Congression. Initially, a bivalent close to one spindle pole becomes mono-oriented and is pulled poleward by flux (blue arrow). Chromosome arms bind to an increasing density of spindle MTs and the resulting resistance (yellow arrows) increases kinetochore tension to induce plus-end polymerization (red arrows). Poleward movement stops, facilitating the capture of the homologous kinetochore by the opposite pole (blue arrows in the spindle). Capture of the unattached kinetochore produces an even greater amount of tension and polymerization (red arrow) at the opposite kinetochore, allowing the bi-oriented bivalent to move to the spindle equator (orange arrow), even though kMT number and flux-generated force (blue arrows) are greater at the lagging homologous kinetochore. Finally, an equilibrium position is established at the spindle equator when the leading kinetochore is captured by an equal number of kMTs. Poleward force from flux (blue arrows) is equivalent in each half-spindle and is balanced by an equal rate of kMT plus-end assembly (red arrows) induced by moderate tension.\u0026lt;\/div\u0026gt;\u0022 data-icon-position=\u0022\u0022 data-hide-link-title=\u00220\u0022\u003E\u003Cspan class=\u0022hw-responsive-img\u0022\u003E\u003Cimg class=\u0022highwire-fragment fragment-image lazyload\u0022 alt=\u0022 Fig. 3. \u0022 src=\u0022data:image\/gif;base64,R0lGODlhAQABAIAAAAAAAP\/\/\/yH5BAEAAAAALAAAAAABAAEAAAIBRAA7\u0022 data-src=\u0022http:\/\/jcs.biologists.org\/content\/joces\/118\/10\/2341\/F3.medium.gif\u0022\/\u003E\u003Cnoscript\u003E\u003Cimg class=\u0022highwire-fragment fragment-image\u0022 alt=\u0022 Fig. 3. \u0022 src=\u0022http:\/\/jcs.biologists.org\/content\/joces\/118\/10\/2341\/F3.medium.gif\u0022\/\u003E\u003C\/noscript\u003E\u003C\/span\u003E\u003C\/a\u003E\u003C\/div\u003E\u003C\/div\u003E\u003Cul class=\u0022highwire-figure-links inline\u0022\u003E\u003Cli class=\u0022download-fig first\u0022\u003E\u003Ca href=\u0022http:\/\/jcs.biologists.org\/content\/joces\/118\/10\/2341\/F3.large.jpg?download=true\u0022 class=\u0022highwire-figure-link highwire-figure-link-download\u0022 title=\u0022Download Fig. 3. \u0022 data-icon-position=\u0022\u0022 data-hide-link-title=\u00220\u0022\u003EDownload figure\u003C\/a\u003E\u003C\/li\u003E\u003Cli class=\u0022new-tab\u0022\u003E\u003Ca href=\u0022http:\/\/jcs.biologists.org\/content\/joces\/118\/10\/2341\/F3.large.jpg\u0022 class=\u0022highwire-figure-link highwire-figure-link-newtab\u0022 target=\u0022_blank\u0022 data-icon-position=\u0022\u0022 data-hide-link-title=\u00220\u0022\u003EOpen in new tab\u003C\/a\u003E\u003C\/li\u003E\u003Cli class=\u0022download-ppt last\u0022\u003E\u003Ca href=\u0022\/highwire\/powerpoint\/1564466\u0022 class=\u0022highwire-figure-link highwire-figure-link-ppt\u0022 data-icon-position=\u0022\u0022 data-hide-link-title=\u00220\u0022\u003EDownload powerpoint\u003C\/a\u003E\u003C\/li\u003E\u003C\/ul\u003E\u003C\/div\u003E\u003Cdiv class=\u0022fig-caption\u0022 xmlns:xhtml=\u0022http:\/\/www.w3.org\/1999\/xhtml\u0022\u003E\u003Cspan class=\u0022fig-label\u0022\u003E\n \u003Cstrong\u003EFig. 3.\u003C\/strong\u003E\n \u003C\/span\u003E \n \u003Cp id=\u0022p-9\u0022\u003EMechanistic models for bivalent positioning and congression in meiotic spindles of grasshopper spermatocytes, using the force from flux and plus-end kMT assembly. (A) Bivalent positioning. A bivalent maintains an equilibrium position at the spindle equator owing to an equivalent amount of flux (blue arrows) on homologous kinetochores attached to an equal number of kMTs. Moderate tension on the kinetochores induces kMT plus-end assembly (red arrows). Laser irradiation (green line) partially destroys a kinetochore, which reduces the number of kMTs to which it can bind. This sudden imbalance in kMT number increases the stress on the remaining kMTs of the irradiated kinetochore, inducing a greater rate of kMT plus-end assembly (red arrow). Compressive force on the opposing kMTs inhibits or decreases the rate of plus-end assembly, and, consequently, the bivalent moves (orange arrow) towards the pole that has the greater number of kMT attachments and the larger flux-generated force (larger blue arrow). As chromosome arms bind to an increasing density of spindle MTs that resist their poleward movement (yellow arrows), the increased tension on the unirradiated kinetochore induces plus-end polymerization. A new equilibrium position is reached when tension-induced plus-end assembly on the unirradiated kinetochore equals the rate of disassembly at the pole. (B) Congression. Initially, a bivalent close to one spindle pole becomes mono-oriented and is pulled poleward by flux (blue arrow). Chromosome arms bind to an increasing density of spindle MTs and the resulting resistance (yellow arrows) increases kinetochore tension to induce plus-end polymerization (red arrows). Poleward movement stops, facilitating the capture of the homologous kinetochore by the opposite pole (blue arrows in the spindle). Capture of the unattached kinetochore produces an even greater amount of tension and polymerization (red arrow) at the opposite kinetochore, allowing the bi-oriented bivalent to move to the spindle equator (orange arrow), even though kMT number and flux-generated force (blue arrows) are greater at the lagging homologous kinetochore. Finally, an equilibrium position is established at the spindle equator when the leading kinetochore is captured by an equal number of kMTs. Poleward force from flux (blue arrows) is equivalent in each half-spindle and is balanced by an equal rate of kMT plus-end assembly (red arrows) induced by moderate tension.\u003C\/p\u003E\n \u003Cdiv class=\u0022sb-div caption-clear\u0022\u003E\u003C\/div\u003E\u003C\/div\u003E\u003Cspan class=\u0022highwire-journal-article-marker-end\u0022\u003E\u003C\/span\u003E\u003C\/div\u003E\u003Cspan id=\u0022related-urls\u0022\u003E\u003C\/span\u003E\u003C\/div\u003E\u003C\/div\u003E\u003C\/div\u003E\u003C\/li\u003E\u003C\/ul\u003E\u003C\/div\u003E\u003C\/div\u003E\u003C\/div\u003E \u003C\/div\u003E\n\n \n \u003C\/div\u003E\n\u003Cdiv class=\u0022panel-separator\u0022\u003E\u003C\/div\u003E\u003Cdiv class=\u0022panel-pane pane-earthchem\u0022 \u003E\n \n \n \n \u003Cdiv class=\u0022pane-content\u0022\u003E\n \u003Ca href=\u0022http:\/\/ecp.iedadata.org\/doidata\/\u0022 class=\u0022\u0022 data-icon-position=\u0022\u0022 data-hide-link-title=\u00220\u0022\u003E\u003Cimg src=\u0022http:\/\/ecp.iedadata.org\/doibanner\/\u0022 alt=\u0022\u0022 \/\u003E\u003C\/a\u003E \u003C\/div\u003E\n\n \n \u003C\/div\u003E\n\u003C\/div\u003E\n \u003C\/div\u003E\n\u003C\/div\u003E\n\u003C\/div\u003E\u003Cscript type=\u0022text\/javascript\u0022 src=\u0022http:\/\/jcs.biologists.org\/sites\/default\/files\/js\/js_hZg96SP9gBcOluDp2mGc57d8sP8uJ7g8P_JYsCISOgQ.js\u0022\u003E\u003C\/script\u003E\n\u003C\/body\u003E\u003C\/html\u003E"}